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NOTES ON GEOGRAPHIC DISTRIBUTION Check List 19 (2): 169-175 


https://doi.org/10.15560/19.2.169 


Occurrence of Pheidole tijucana Borgmeier, 1927 (Hymenoptera, 
Formicidae) in the state of Sao Paulo, Brazil 


DEBORA CRISTINA DE OLIVEIRA GONGALVES', LivIA PIRES DO PRADO’, FABRICIO SEVERO MaG- 
ALHAEsS!, MARLINA RIBEIRO ABONIZIO-SANTOS!, MARIA SANTINA DE CASTRO Morini" 


1 Laboratério de Mirmecologia do Alto Tieté, Nucleo de Ciéncias Ambientais, Nucleo de Ciéncias Ambientais, Universidade Mogi das 
Cruzes, Mogi das Cruzes, SP, Brazil e DCOG: debora_cog@outlook.com @ https://orcid.org/0000-0002-3739-9786 e FSM: fabricio- 
severo.fa@gmail.com https://orcid.org/0000-0002-7173-658X e MRAS: marlinaribeiro@live.com ® https://orcid.org/0000-0002- 
6637-7236 e MSCM: mscmorini@gmail.com ® https://orcid.org/0000-0002-1823-6703 

2 Laboratério de Hymenoptera, Museu de Zoologia da Universidade de Sao Paulo, Sao Paulo, SP, Brazil « livia.pires7@gmail.com 
https://orcid.org/0000-0003-1819-8767 

* Corresponding author 


Abstract. Pheidole Westwood, 1839 is a hyperdiverse ant genus that is widely distributed in a variety of terres- 
trial ecosystems, especially in the tropics. The identification of Pheidole species is challenging, thus contributing 
to the description of several questionable morphospecies in the literature and making the geographic distribution 
of most species uncertain. In this work, we present the first records of Pheidole tijucana Borgmeier, 1927 from the 
state of Sao Paulo, Brazil. The specimens were recorded at Parque Ibirapuera Conservacao, an urban green area 
located in the southern part of the city of Sao Paulo. The sampling was performed in 2015, 2016, and 2017 using 
pitfall traps. In total, 224 specimens (one major worker and 223 minor workers) of P. tijucana were recorded. Our 
new data help fill a distribution gap and highlights the importance of reporting species that are a challenge to 
identify. 


Keywords. Atlantic Forest, biodiversity, Myrmicinae, urban green area, urban park 


Academic editor: Sandor Buys 
Received 25 January 2023, accepted 5 March 2023, published 14 March 2022 


Gongalves DCO, Prado LP, Magalhaes FS, Abonizio-Santos MR, Morini MSC (2023) Occurrence of Pheidole tijucana Borgmeier, 1927 
(Hymenoptera, Formicidae) in the state of Sao Paulo, Brazil. Check List 19 (2): 169-175. https://doi.org/10.15560/19.2.169 


Introduction 


Pheidole Westwood, 1839 (Myrmicinae) is a hyperdi- 
verse ant genus with more than 1,000 valid species and 
subspecies, which are widely distributed around the 
world. In the Americas, Pheidole is the richest genus of 
Myrmicinae (Kempf 1972; Wilson 1976), having been 
recorded in the most varied environmental conditions 
and habitats (Economo 2015), in disturbed (Jesovnik 
and Bujan 2021), preserved (Jahan 2022), and regenera- 
ting areas (Aguiar 2022). Around 700 species are found 
in the Neotropical Region, 200 of which occur in Brazil 
and 60 of which occur in the state of Sao Paulo (Janicki 
et al. 2016). 

Pheidole species display morphological and behav- 


ioral traits that enabled a great adaptive success (i.e., the 
division of labor in two morphologically distinct castes; 
loss of a functional sting; the complete absence of ova- 
ries in the worker castes) (Holldobler and Wilson 1990; 
Wilson 2003). Nesting and foraging occur mainly in 
deeper soil layers, or above or between the interstices 
of the leaf litter (Delabie et al. 2000; Casadei-Ferreira 
2021). In the leaf litter, Pheidole may occupy fruits, 
seeds, and wood in varying degrees of decomposition 
(Castaho-Menzes et al. 2015; Casadei-Ferreira 2021), 
and the latter includes twigs (Fernandes et al. 2019). 
Barroso et al. (2020) and Almeida et al. (2021) suggest- 
ed that the occupation of twigs by species of Pheidole 
could be due to a process of selection. 

Pheidole tijucana Borgmeier, 1927 belongs to the 


©The authors. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, 
distribution, and reproduction in any medium, provided the original author and source are credited. 


170 


fallax group (Wilson 2003), although its biology 
remains unknown. The species was described in 1927 
by Borgmeier, based on 15 major workers and 12 minor 
workers collected by him in Tijuca, a neighborhood in 
the city of Rio de Janeiro (Rio de Janeiro state, Brazil). 
Many decades later, this species was again recorded, 
this time from the state of Bahia (Santos et al. 2017), in 
remnants of the Atlantic Forest at different regenera- 
tion stages; Santos et al. used pitfall traps and Winkler 
extractors to collect the specimens. Although Sao Pau- 
lo is one of the Brazilian states best-sampled for its ant 
fauna, no other records of P. tijucana have been record- 
ed since, even in the recent updates on the Atlantic For- 
est species (Feitosa et al. 2021; Silva et al. 2022). Given 
the scarcity of information on P. tijucana, we report the 
first records of this species from the state of Sao Paulo. 


Methods 


The specimens were collected in an urban green area, 
Parque Ibirapuera Conservacao, which is in the city 
of Sao Paulo. The park is within the Brazilian Atlan- 
tic Forest domain (Ribeiro et al. 2009); it has an area 
of 1,584 m’ and is composed of mixed woodland with 
Eucalyptus and other exotic species, as well as native 
species. There are flower beds, lawns, and theme gar- 
dens on site, with a total of 532 species of vascular plan- 
ts, including trees, shrubs, and herbaceous plants. The 
climate is humid subtropical, characterized by a noto- 
riously dry winter and a rainy summer; temperatures 


23°34,920'S 


Sao Paulo 


23°35,340'S 


© Distribution of Pheidole tiucana 
@ New record of Pheidole tijucana 
M) Atlantic Forest 


46°39,960'W 
- ae Se 


Check List 19 (2) 


vary between 12 °C during the winter and 28 °C during 
the summer (annual average temperature 20.3 °C). 

Our expeditions to collect specimens were carried 
out in the summers and winters of 2015, 2016, and 2017 
in four areas of the park (Fig. 1). Ants were collected 
with pitfall traps containing 500 mL of preserving liq- 
uid (70% ethanol and 10% formaldehyde); the traps 
were buried at ground level. In total, 100 pitfall traps 
were used, with each area having 25 traps. Each trap 
was placed 3 m away from the next closest trap. The 
traps remained on site for eight days. 

The genus was identified using the key by Baccaro 
et al. (2015). The species identification used the keys by 
Wilson (2003). The specimens collected were also com- 
pared to the type specimens of P. tijucana, which are 
deposited at Cole¢ao de Hymenoptera of the Museu 
de Zoologia da Universidade de Sao Paulo (MZSP). 
The vouchers were also deposited at MZSP (individu- 
ally as MZSP67490, MZSP67491, MZSP67492, and 
MZSP95643) and at the Laboratorio de Mirmecolo- 
gia do Alto Tieté da Universidade de Mogi das Cruzes 
(LAMAT) (Souza-Campana et al. 2020). 


Results 


Pheidole tijucana Borgmeier, 1927 
Figures:2,3 


New records. BRAZIL - SAo PAULO « Sao Paulo, Par- 
que Ibirapuera Conservacao; 23°35'38"N, 046°39'52" W; 


S0PE'SEoES S,0Z6 E087 


S,092'SEoET 


46°39,240'W 


Figure 1. Geographic location of the areas of occurrence of Pheidole tijucana in the Brazilian Atlantic Forest. A. Study site, Sao 
Paulo, Brazil. B. Parque Ibirapuera Conservagao, in the city of Sao Paulo. 


Gongalves et al. | Pheidole tijucana in the state of Sao Paulo 


171 


1mm 


Figure 2. Minor worker of Pheidole tijucana. A. Head in frontal view. B. Latero-dorsal view. C. Lateral view. 


761 alt.; 29.V.2015; R. Bertani col. leg.; pitfall; 569, 
MZSP; 1 major worker, MZSP95643 « same data, except 
23°35'33"N, 046°39'44"W; 757 alt.; 22.V.2015; R Ber- 
tani col. leg.; pitfall; 49, LAMAT « same data, except 
23°35'38"N, 046°39'52"W; 761 alt.; 29.1.2016; R. Ber- 
tani col. leg.; pitfall; 108 minor workers, LAMAT; 19, 
MZSP67490; 1 minor worker, MZSP67491; 1 minor 
worker, MZSP67492 « same data, except 23°35'33"N, 
046°39'44"W; 757 alt.; 29.1.2016; R Bertani col. leg.; pit- 
fall; 13 minor workers, LAMAT «¢ same data, except 
23°35'28"N, 046°39'19"W; 765 alt.; 29.1.2016; R Bertani 
col. leg.; pitfall; 1 minor workers, LAMAT e same data, 
except 23°35'38"N, 046°39'52"W; 761 alt.; 14.IX.2016; 


R. Bertani col. leg.; pitfall; 16 minor workers, LAMAT 
e same data, except 23°35'38"N, 046°39'52" W; 761 alt.; 
14.11.2017; R Bertani col. leg.; pitfall; 19 minor workers, 
LAMAT « same data, except 23°35'33"N, 046°39'44" W; 
757 alt.; 14.11.2017; R Bertani col. leg.; pitfall; 1 minor 
worker, LAMAT « same data, except 23°35'28"N, 046° 
39'19"W; 765 alt.; 14.11.2017, R Bertani col. leg.; pitfall; 2 
minor worker, LAMAT. 

Total number of specimens: 223 minor workers and 
1 major worker. 


Identification. As reported by Wilson (2003), the minor 
worker of P. tijucana can be identified according to 


172 


Check List 19 (2) 


Figure 3. Major worker of Pheidole tijucana. A. Head in frontal view. B. Lateral view. C. Dorsal view. 


the morphological characteristics of the fallax species 
group; also, it is distinct from all other species in this 
group because of the following characteristics: occiput 
constricted to the neck a with broad nuchal collar (Fig. 
2A, B), propodeal spines reduced to denticles (Fig. 2C), 
and light brown, except for the head, which is a darker, 


reddish shade (Fig. 2). The major worker (Fig. 3A-C), 
as reported by Wilson (2003), is morphologically simi- 
lar to Pheidole lucretii Wilson 2003. However, the two 
species are distinguished mainly by the sculpture on 
the surface of the head. In P. tijucana most of the occi- 
put vertex and frontal lobes are smooth and shiny (Fig. 


Gongalves et al. | Pheidole tijucana in the state of Sao Paulo 


3A, red arrow), with a presence of short longitudinal 
carinulae around midline at the occiput (Fig. 3A, blue 
arrow), whereas in P. lucreti the same region of the head 
is superficially rugoreticulate. 


Discussion 


Pheidole tijucana was found in an urban green area 
inserted in the Brazilian Atlantic Forest domain. Our 
results, as for all previous records of this species (Borg- 
meier 1927; Santos et al. 2017), suggest that it inhabits 
dense tropical rainforests. The Atlantic Forest consists of 
a mosaic of vegetation with distinct forests. One of them 
is the dense rainforest, which is characterized by very 
high humidity, high temperatures (average of 25 °C), and 
high precipitation well distributed throughout the year 
(0-60 dry days); vegetation is composed of phanerophy- 
tes, lianas, and epiphytes (IBGE 2012). This phytophy- 
siognomy presents a wide diversity of ants (Feitosa et al. 
2021; Silva et al. 2022), given that in 1 m’ of leaf litter it is 
possible to find up to 35 species (Silva and Brandao 2010). 

Since the description of P. tijucana, many surveys of 
leaf litter have been carried out in the Brazilian Atlan- 
tic Forest (i.e., Delabie et al. 2007; Pacheco et al. 2009; 
Suguituru et al. 2011, 2013; Silva et al. 2022), includ- 
ing in wood structures that are colonized by species 
of Pheidole (Souza et al. 2012; Fernandes et al. 2019, 
2020). Moreover, several surveys have also been made 
in urban areas of Brazil (Bueno et al. 2017), and P. tiju- 
cana has not been recorded in any of them. The scar- 
city of records for P. tijucana may indicate that the 
species is not abundant and has a more limited dis- 
tribution. However, the low taxonomic resolution of 
hyperdiverse genera such as Pheidole should be taken 
into consideration (Kempf 1961; Silva et al. 2022). Col- 
onies of the Pheidole species typically have hundreds 
to thousands of workers (minor and major), but here 
we sampled only one major worker in our sampling, 
which may reflect the collection method used. In gen- 
eral, minor workers are usually responsible for forag- 
ing, while major workers remain mainly inside the nest 
(e.g., Wetterer, 2007). Thus, it is possible that the spe- 
cies was simply not identified in the above-mentioned 
checklists, and that its distribution and abundance are 
actually higher than what is now thought. 

The lack of knowledge on the distribution and biol- 
ogy of insect species is alarming, mainly in face of the 
decline in the populations of these invertebrates (San- 
chez-Bayo and Wyckhuys 2019; Wagner 2020). Unfor- 
tunately, our comprehension of tropical biodiversity is 
still severely limited by significant deficiencies in our 
knowledge of the taxonomy and distribution of species 
(Hortal et al. 2015). Hence, the occurrence record of P. 
tijucana in the state of Sao Paulo, in an urban area of a 
huge metropolis is quite surprising, and it opens new 
possibilities for the study of its biology. 


Acknowledgements 
We thank the National Council for Scientific and Tech- 


173 


nological Development (CN Pq, process no. 126691/2021- 
5) for the undergraduate research fellowship granted to 
the first author. L.P. Prado acknowledges the Sao Pau- 
lo Research Foundation (FAPESP) for the postdoctoral 
fellowship (grant no. 2022/01974-8). 


Author Contributions 


Conceptualization: DCOG, LPP, MSCM. Data cura- 
tion: DCOG, FSM, MRAS. Formal analysis: DCOG, 
LPP. Investigation: DCOG, FSM, MRAS, MSCM, LPP. 
Methodology: LPP. Project administration: DCOG. 
Supervision: MSCM. Writing - original draft: DCOG. 
Writing - review and editing: DCOG, FSM, MRAS, 
MSCM, LPP. 


References 


Aguiar JJM, Anjos DV, Carvalho RL, Almeida RW, Santos 
ACC, Santos JC (2022) Plant richness drives ant diversity 
in Eucalyptus-dominated landscape on Brazilian savanna. 
Austral Ecology 47 (1): 17-25. https://doi.org/10.1111/ 
aec. 13060 

Almeida RPS, Andrade-Silva J, Silva RR, Fernandes TT, 
Souza-Campana DR, Silva NS, Morini MSC (2021) 
Twigs in leaf litter: overlap in twig size occupation by 
nesting ants. Insectes Sociaux 68 (2): 199-206. https://doi. 
org/10.1007/s00040-021-00816-6 

Baccaro FB, Feitosa RM, Fernandez F, Fernandes IO, Izzo 
TJ, Souza JD, Solar R (2015) Guia para os géneros de for- 
migas do Brasil. Editora INPA, Manaus, Brazil, 388 pp. 

Barroso SC, Longui EL, Fernandes TT, Oliveira CM, Fer- 
reira AC, Silva RR, Morini MSC (2020) Twigs occupied 
by Pheidole Westwood, 1839: is there a difference between 
species?. Biota Neotropica 20 (2): e20190897. https://doi. 
org/10.1590/1676-0611-BN-2019-0897 

Borgmeier T (1927) Algumas novas formigas brasileiras. 
Arquivos do Museu Nacional 29: 57-65. 

Bueno OC, de Carvalho Campos AE, Morini MSC (2017) 
Formigas em ambientes urbanos no Brasil. 6 ed. Canal, 
Bauru, Brazil, 685 pp. 

Casadei-Ferreira A (2021) Pheidole. In: Starr CK (Ed.) Ency- 
clopedia of social insects. Springer, Cham, Switzerland, 
728-731. https://doi.org/10.1007/978-3-030-28102-1 

Castano-Meneses G, Mariano CF, Rocha P, Melo T, Tavar- 
es B, Almeida E, Silva LD, Pereira TP, Delabie JH (2015) 
The ant community and their accompanying arthropods 
in cacao dry pods: an unexplored diverse habitat. Duge- 
siana 22 (1): 29-35. https://doi.org/10.32870/dugesiana. 
v22i1.4173 

Delabie JH, Jahyny B, Nascimento IC, Mariano CS, Lacau 
S, Campiolo S, Philpott SM, Leponce M (2007) Contri- 
bution of cocoa plantations to the conservation of native 
ants (Insecta: Hymenoptera: Formicidae) with a special 
emphasis on the Atlantic Forest fauna of southern Bahia, 
Brazil. Biodiversity and Conservation 16 (8): 2359-2384. 
https://doi.org/10.1007/s10531-007-9190-6 

Delabie JHC, Agosti D, Nascimento IC (2000) Litter ant 
communities of the Brazilian Atlantic rain forest region. 
In: Agosti D, Majer J, Alonso L, Schultz T (Eds.) Sampling 


174 


ground-dwelling ants: case studies from the world’s rain 
forests. School of Environmental Biology, Bulletin 18: 
1-17. 

Economo EP, Klimov P, Sarnat EM, Guénard B, Weiser 
MD, Lecroq B, Knowles LL (2015) Global phylogenetic 
structure of the hyperdiverse ant genus Pheidole reveals 
the repeated evolution of macroecological patterns. Pro- 
ceedings of the Royal Society B: Biological Sciences 282 
(1798): 20141416. https://doi.org/10.1098/rspb.2014.1416 

Feitosa RM, Morini MSC, Martins AC, Ribeiro TM, Noll 
FB, Santos EK, Cancello EM, Constantini JP (2021) 
Social insects of the Atlantic Forest. In: Marques MCM, 
Grelle CEV (Eds.) The Atlantic Forest. Springer, Cham, 
Switzerland, 151-183. https://doi.org/10.1007/978-3- 
030-55322-7 

Fernandes TT, Dattilo W, Silva RR, Luna P, Braz AB, 
Morini MSC (2020) Cohabitation and niche overlap in 
the occupation of twigs by arthropods in the leaf litter of 
Brazilian Atlantic Forest. Insectes Sociaux 67 (2): 239-47. 
https://doi.org/10.1007/s00040-020-00753-w 

Fernandes TT, Dattilo W, Silva RR, Luna P, Oliveira CM, 
Morini MSC (2019) Ant occupation of twigs in the 
leaf litter of the Atlantic Forest: influence of the envi- 
ronment and external twig structure. Tropical Con- 
servation Science 12: 1940082919852943. https://doi. 
org/10.1177/1940082919852943 

H6lldobler B, Wilson EO (1990) The ants. Belknap Press of 
Harvard University Press, Cambridge, USA, 732 pp. 

Hortal J, Bello F, Diniz-Filho JAF, Lewinsohn TM, Lobo 
JM, Ladle RJ (2015) Seven shortfalls that beset large- 
scale knowledge of biodiversity. Annual Review of Ecol- 
ogy, Evolution and Systematics 46: 523-549. https://doi. 
org/10.1146/annurev-ecolsys-112414-054400 

IBGE (Instituto Brasileiro de Geografia e Estatistica) (2012) 
Manual técnico da vegetacao brasileira: sistema fitogeo- 
grafico, inventario das formacoes florestais e campestres, 
técnicas e manejo de colegées botanicas, procedimentos 
para mapeamentos. 2nd edition. Coordenacao de Recur- 
sos Naturais e Estudos Ambientais, Rio de Janeiro, Brasil, 
276 pp. 

Jahan MN, Rahman MM, Miah MG, Ahamed T (2022) 
Effects of vegetation types and habitat disturbance on 
species richness and form composition of ants (Hyme- 
noptera, Formicidae) in Lawachara National Park, Bang- 
ladesh. Environment and Ecosystem Science 6 (1): 01-06. 
http://doi.org/10.26480/ees.01.2022.01.06 

Janicki J, Narula N, Ziegler M, Guénard B, Economo EP 
(2016) Visualizing and interacting with large-volume 
biodiversity data using client-server web-mapping appli- 
cations: The design and implementation of antmaps. 
org. Ecological Informatics 32: 185-193. https://doi. 
org/10.1016/j.ecoinf.2016.02.006 

JeSovnik A, Bujan J (2021) Wooded areas promote species 
richness in urban parks. Urban Ecosystems 24 (6): 1305- 
1315. https://doi.org/10.1007/s11252-021-01112-8 

Kempf WW (1961) A survey of the ants of the soil fauna in 
Surinam (Hymenoptera: Formicidae). Studia Entomolo- 
gica 4: 81-524. 

Kempf WW (1972) Catalogo abreviado das formigas da 
regiao neotropical (Hymenoptera: Formicidae). Studia 


Check List 19 (2) 


Entomologia 15: 1-344. 

Pacheco R, Silva RR, Morini MSC, Brandao CR (2009) 
A comparison of the leaf-litter ant fauna in a secondary 
atlantic forest with an adjacent pine plantation in south- 
eastern Brazil. Neotropical Entomology 38 (1): 55-65. 
https://doi.org/10.1590/S1519-566X2009000100005 

Ribeiro MC, Metzger JP, Martensen AC, Ponzoni FJ, Hiro- 
ta MM (2009) The Brazilian Atlantic Forest: How much is 
left, and how is the remaining forest distributed? Implica- 
tions for conservation. Biological Conservation 142 (6): 
1141-1153. https://doi.org/10.1016/j.biocon.2009.02.021 

Sanchez-Bayo F, Wyckhuys KA (2019) Worldwide decline 
of the entomofauna: a review of its drivers. Biologi- 
cal conservation 232: 8-27. https://doi-org/10.1016/j. 
biocon.2019.01.020 

Santos RJ, Koch EBA, Leite CMP, Porto TJ, Delabie JHC 
(2017) An assessment of leaf-litter and epigaeic ants 
(Hymenoptera: Formicidae) living in different landscapes 
of the Atlantic Forest Biome in the state of Bahia, Bra- 
zil. Journal of Insect Biodiversity 5 (19): 1-19. https://doi. 
org/10.12976/jib/2017.5.19 

Silva RR, Brandao CRF (2010) Morphological patterns and 
community organization in leaf-litter ant assemblag- 
es. Ecological Monographs 80 (1): 107-124. https://doi. 
org/10.1890/08-1298.1 

Silva RR, Martello FE, Feitosa RM, Silva OGM, Prado LP, 
Brandao CR, Albuquerque EZ, Morini MSC, Delabie 
JH, Monteiro ECS, Ribeiro MC (2022) Atlantic ants: a 
data set of ants in Atlantic Forests of South America. Eco- 
logy 103: e03580. https://doi.org/10.1002/ecy.3580 

Souza-Campana DR, Wazema CT, Magalhaes FS, Sil- 
va NS, Nagatani VH, Suguituru SS, Goto MA, Mori- 
ni MSC (2020) Colecao de referéncia do Laboratério 
de Mirmecologia do Alto Tieté, Sao Paulo, Brasil: status 
atual e perspectivas. Boletim do Museu Paraense Emilio 
Goeldi-Ciéncias Naturais 15 (1): 317-336. https://doi. 
org/10.46357/bcnaturais.v15i1.274 

Souza DR, Fernandes TT, Nascimento JR, Suguituru SS, 
Morini MSC (2012) Characterization of ant communi- 
ties (Hymenoptera: Formicidae) in twigs in the leaf lit- 
ter of the Atlantic rainforest and eucalyptus trees in the 
southeast region of Brazil. Psyche: a Journal of Entomolo- 
gy 2012: 1-12. https://doi.org/10.1155/2012/532768 

Suguituru SS, Silva RR, Souza DRD, Munhae CDB, Morini 
MSC (2011) Ant community richness and composition 
across a gradient from Eucalyptus plantations to second- 
ary Atlantic Forest. Biota Neotropica 11 (1): 369-376. 
https://doi.org/10.1590/S1676-06032011000100034 

Suguituru SS, Souza DR, Munhae CD, Pacheco R, Morini 
MSC (2013) Ant species richness and diversity (Hyme- 
noptera: Formicidae) in Atlantic Forest remnants in the 
Upper Tieté river basin. Biota Neotropica 13: 141-152. 
https://doi.org/10.1590/S1676-06032013000200013 

Wagner DL (2020) Insect declines in the Anthropocene. 
Annual Review of Entomology 65: 457-480. https://doi. 
org/10.1146/annurev-ento-011019-025151 

Wetterer JK (2007) Biology and impacts of Pacific Island 
invasive species. 3. The African big-headed ant, Pheidole 
megacephala (Hymenoptera: Formicidae) Pacific Sci- 
ence 61 (4): 437-456. https://doi.org/10.2984/1534-6188 


Gongalves et al. | Pheidole tijucana in the state of Sao Paulo 


(2007)61[437:baiopi]2.0.co;2 

Wilson EO (1976) Which are the most prevalent ant genera. 
Studia Entomologica 19 (1-4): 187-200. 

Wilson EO (2003) Pheidole in the New World: a dominant, 
hyperdiverse ant genus. Harvard University Press, Cam- 
bridge, USA, 794 pp. 


175 


Data Resources 


The data underpinning the analysis reported in this 
paper are deposited in the Global Biodiversity Infor- 
mation Facility and are available at https://doi.org/ 
10.15468/rv7tdk.